Bulletin of the
British Museum (Natural History)

The songs of the western European
grasshoppers of the genus Stenobothrus in
relation to their taxonomy (Orthoptera:
Acrididae)

D. R. Ragge

Entomology series
Vol55 No2 24 September 1987

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ISBN 0 565 06032 5
ISSN 0524-6431 Entomology series
Vol 55 No 2 pp 393-424
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 24 September 1987

The songs of the western European grasshoppers of
the genus Stenobothrus in relation to their taxonomy
(Orthoptera: Acrididae)

D. R. Ragge

Department of Entomology, British Museum (Natural History), Cromwell Road, London
SW7 5BD

Contents
PERI N FY SUR oe iyi so ets ctrrache senting eke Aaa MAA sate Re Mahe wiynwsaudvleavenivedinndndavanenniisan 393
BEYETEMCAUCILORN Soca Sane coterie tear adeaaen eacie.s diee en aaah Anions onis cl calsidorawseas'a cabin in aninyscanied v/ansia ter 393
PA CKMOW I COPEMEM (Stems onrme tate me ee ntee iors etter entre cea tania cs <rdcsertesmeseaessnaiesi 394
PX DOLE VIA TONS OL CC POSIOGIES caarersiicn: states roe eosseber sess sedeasecdeesatervosesadedonsetescese se 394
SARI RERIGR A testa er, 2o J nccPe Nc gute est ace sce ever sT AREY « cunadee Jan un vende due sin stlia en aabuavad deawsvati« 395
RECOLGIN gan dlanalySiie tHe SOMPSs oat ns-ceeee see het adaceewasns apkieua= Wa nateccnoanenctestenes 395
SON CMETOMMOlO CViternne ee Nen cere nce actetac te tease ane eca re et ne eetnanttinsione eeting awake eansehey 397
Rea Ms ANON roe crn's duty tein as seh ecideicla at nF <-ig ss AN deaets Sniaithas Zara cello «Shak Mh dade a disawadiesdesiea 397
SUDESON rasa TAN ONOMIC CHALACTEN 5 ss. cevac aoadatine det aeraedes eye «clewe atsaleatendiatideee cra visiee ston’ 398
BP Lene MENTE CNEL MINS st sac cue oi meen cscie vimev'nnechigesifhanenwesnssantnndnvabamainddnceenedesan 399
RRP RRIENE SR RINE MER Mae esto coh on xn vw 2c Aaidy athitan Sener asad Gam cnmpewadnemeniien'nhinnns 399
Keys to the principal western European species of Stenobothrus ...............0cc0cceeeeeee 400
Key based pronarily ommorpholopical ChatacCters:..........0.c-se.scerssscccavcucaseeravenses 400
PREY ASE OM SOME AR ARL CES rien coca scuaviansccecesternedswosenwasheseseasewaseoaracavennecersanse 402
The songs of the westerm European StenODOMris «6s... cccssseccedseseccenensaesenecsscsecees 404
LNGHSTIRETT TSS Gor aneuioc dene dncoocee RO GusBagE CEA nUCUPRTP aoa cepcoCadnee Te ntaaene CaCC ee EE BHP eT CEE RE anE EEE 422
NCEE CORN tes eis ecg at Wh sspint one eens Maar lesb aOR nae cath Uacasmcwdsiiiadnonvadedvemecoseradsas 424

Synopsis

The male calling songs of nine species of the Gomphocerine genus Stenobothrus are described and fully
illustrated with oscillograms at several different speeds. The courtship song is also described for the seven
species in which such a song is known. The songs of three species (festivus, bolivarii and apenninus) are
described for the first time. For each species a list is provided of references to any past published work on
the songs, classified according to the kind of information given. Notes on recognition, using both
morphology and song, are given for each species, and two identification keys are provided, one based
mainly on morphology and the other on song. The value of the song as a taxonomic character is discussed.

Introduction

Stenobothrus is one of the better known genera of Gomphocerinae, a group that includes most of
the common European grasshoppers of open grassland and moorland. They are well known for
their characteristic songs and are widely used in bio-acoustic, ecological and cytogenetic
research. The subfamily includes about 150 European species, of which over 20 are pests in parts
of their range, seven of them being regarded as major ones (Tsyplenkov, 1970).

Stenobothrus is a widespread Palaearctic genus, including over 30 species living in habitats
ranging from lush grassland to steppe and high mountains. Eleven species have been recorded
from western Europe, of which three, /ineatus, nigromaculatus and fischeri, become pests when
they occur in sufficient numbers, causing damage to cereal grasses, hayfields and alfalfa
(Bei-Bienko & Mishchenko, 1951; Tsyplenkov, 1970).

It is now well known that the songs of Gomphocerine grasshoppers are of great taxonomic
value. Their role as a mate recognition system makes them particularly valuable in resolving

Bull. Br. Mus. nat. Hist. (Ent.) 55 (2): 393-424 Issued 24 September 1987

394 D. R. RAGGE

complexes of sibling species and in establishing the status of local populations showing small
morphological differences. There has so far been no comprehensive account of these songs,
presented group by group and illustrated with oscillograms. The main aim of the present paper,
and of similar papers already published (Ragge, 1986) or in preparation, is to provide such an
account for the western European species, arranged systematically by group and fully illustrated
with oscillograms at several different speeds.

No information has previously been published on the songs of three of the nine species
included in this study (festivus, bolivarii and apenninus) and no oscillograms have previously
been published for a further two (fischeri and grammicus). For the species in which published
information on the song is already available, references to the sources are given, classified
according to the kind of information published.

Acknowledgements

I am much indebted to Drs A. Gorochov and A. Kaltenbach, who have kindly lent me
type-specimens or other material from their respective institutions. I should also like to thank
Mr C. F. Dewhurst and Miss T. M. Howard, who kindly brought me live males of S. stigmaticus
from the Isle of Man, and Mr E. C. M. Haes, who was kind enough to give me a live male of S.
rubicundulus that he had collected in the Italian Alps.

I am particularly grateful to my colleague Mr W. J. Reynolds, who made the studio recordings
used in this study, helped me in reviewing the past literature on the songs, tested the
identification key based on morphological characters and provided valuable comments at
various stages in the preparation of the paper.

Finally I wish to thank my wife, who has given me invaluable help in all my field-work.

Abbreviations of depositories

BMNH _ British Museum (Natural History), London
MNCN Museo Nacional de Ciencias Naturales, Madrid

NM Naturhistorisches Museum, Vienna
ZI Zoological Institute of the Academy of Sciences of the U.S.S.R., Leningrad
Methods

Recording and analysing the songs

All the field recordings of the songs used in this study were made in full sunshine (except
recording 184/1 — see p. 405) using a Uher 4000, 4200 or 4200IC tape recorder and AKG D202
microphone; the tape speed was always 19 cm/s. The subjects were approached stealthily so that
the microphone could be held (or rested on the ground) about 10 cm from them without causing
any disturbance.

The studio recordings were made in the BMNH Acoustic Laboratory using a Kudelski Nagra
IV tape recorder and Sennheiser MKH405 microphone or (recordings 119 and 121 only)
Kudelski Nagra III tape recorder and AKG C61 microphone; the tape speed was 19 or 38 cm/s.
In every case a bench lamp was used to provide light and radiant heat.

Further data are given in Tables 1 and 2 for the recordings used for the oscillograms
reproduced in Figs 9-116. However, the song descriptions given for each species are in most
cases based on many more recordings of numerous songs, all of which were analysed oscillo-
graphically. The oscillograms reproduced in the figures were chosen as being typical of the
species concerned and, in some cases, showing the extent of intraspecific variation. Although I
have given in Tables 1 and 2 the ambient air temperatures at the time of recording, it is really the
body temperature of the singing insect that is important, and this is more dependent on radiant
heat than on the air temperature; as the insects were always receiving radiant heat from the sun
or a bench lamp, the conditions under which the recordings were made were more uniform than
the different air temperatures might suggest.

All the oscillograms were made from recorded songs with a Mingograf 34T ink-jet recorder.

SONGS OF THE WESTERN EUROPEAN Stenobothrus 395

Table 1 Data for the field recordings of male songs of Stenobothrus used for oscillograms reproduced in
this study. All these recordings were made from different males. Recordists’ names are abbreviated as
follows: DRR = D. R. Ragge; WJR = W. J. Reynolds.

Date recorded
and collected

Shade air Recording

Species Locality Recordist temperature No.

lineatus

nigromaculatus

fischeri

festivus

grammicus

bolivarii

stigmaticus

FRANCE: Seine-
Maritime, near
Gamaches

Iva.y: Lazio, Monti
Reatini, foot of
chair-lift to Monte
Terminilletto, 1700 m

FRANCE: Pyrénées-
Orientales, near
Saillagouse, Val d’Eyne

FRANCE: Lozére, Causse
Méjean, near
Crosgarnon

Same locality

Itay: Abruzzi, Gran
Sasso d'Italia, Fossa
di Paganica, 1680 m

ITta.y: Marches, Monti
Sibillini, Forca di
Presta, 1540 m

FRANCE: Vaucluse, Mont
Ventoux, road from
Bédoin, 1350 m

Same locality

FRANCE: Alpes-
Maritimes, near Col
de Vence, 930 m

SPAIN: Madrid, near

Guadarrama, Embalse de

la Jarosa

FRANCE: Vaucluse, Mont
Ventoux, near Pavillon
de Rolland, 800 m

FRANCE: Vaucluse, Mont
Ventoux, road from
Bédoin, 1100 m

SPAIN: Madrid, near
Navacerrada, Embalse
de Navalmedio, 1300 m

FRANCE: Haute-Vienne,
near St
Yrieix-la-Perche

SPAIN: Madrid, near
Navacerrada, Valle de
Barranca

11. viii. 1983

11.ix.1985

22.1x.1976

31.vii. 1982
31.vii.1982

31. viii. 1985

14.ix.1985

15. viii. 1977
15. viii. 1977

25.vili.1977

19.vi.1984

11.viii.1977

12.vili.1977

26. vii. 1983

30.vili.1978

27.vii. 1983

DRR

DRR

WJR

DRR
DRR

DRR

DRR

DRR
DRR

DRR

DRR

DRR

DRR

DRR

WJR

DRR

24°C

18°C

16°C

16°C
16°C

23°C

18°C

2a
2a°€

23 ©

22 C

2356

Lee

26°C

20°C

22°C

511/2

608/4

184/1

482/2
482/3

603/4

609/10

237/1
237/3

241/2

531/6

233/4

234/1

501/8

268/2

503/6

396 D. R. RAGGE

Table 1 — cont.

Data recorded Shade air Recording

Species Locality and collected Recordist temperature No.
apenninus Iraty: Abruzzi, Gran

Sasso d’Italia, Fossa

di Paganica, 1680 m 31.viii.1985 DRR 24°C 603/2

Same locality 31.vili.1985 DRR 19°@ 603/3

ITvAcy: Lazio, Monti

Reatini, foot of

chair-lift to Monte

Terminilletto, 1700 m 11.ix.1985 DRR 18°C 608/1

rubicundulus FRANCE: Alpes-
Maritimes, near St
Dalmas de Tende, La
Miniére de Vallaure,
1500 m 24. vili.1977 DRR 23°C 240/4

ITALY: Trentino-Alto

Adige, near Riva del

Garda, Tremalzo,

1200 m 27. viii. 1985 DRR 13°C 601/5

Table2 Data for the studio recordings of male songs of Stenobothrus used for oscillograms reproduced in
this study. All these recordings were made from different males except for 369 and 371, which were made
on different days from the same male of rubicundulus. The recordist was W. J. Reynolds in every case.

Date Date Air Recording

Species Locality collected recorded temperature No.
lineatus ENGLAND: Kent,

near Wrotham,

Trottiscliffe 10. viii. 1978 11.vili.1978 PEAS 260

Same locality 12.viii.1978 13.vili.1978 29°C 321/1

Same locality 12.viii.1978 13.vili.1978 28°C 322/2
nigromaculatus FRANCE: Pyrénées-

Orientales, near

Saillagouse, Val

d’Eyne 22.ix.1976 29.ix.1976 Zi-€ 176
festivus SPAIN: Madrid, near

Guadarrama, Embalse

de la Jarosa 19. vi.1984 4.vii.1984 — S27
stigmaticus ISLE OF MAN:

Langness Peninsula 1-6.viii.1974 8.viti.1974 24°C 119

Same locality 1-6.vili.1974 = 9. vii. 1974 24°C 121

Same locality 30. viii. 1983 6.i1x.1983 23°C 497
rubicundulus ITALY: Belluno,

Pordoi Pass, below
cable line to
Pordoi Joch 4.1x.1980 16.ix.1980 26.6 369

Same locality 4.ix.1980 18.1x.1980 25°C 371

SONGS OF THE WESTERN EUROPEAN Stfenobothrus 397
Song terminology

The bio-acoustic terms used are defined as follows.
Calling song. The song produced by an isolated male.
Courtship song. The special song produced by a male when close to a female.
Syllable. The sound produced by one complete up and down movement of the hind legs
(Fig. 1).
Echeme. A first-order assemblage of syllables (Fig. 1).
Echeme-sequence. A first-order assemblage of echemes (Fig. 1).
Momentary breaks in the sound (of at least 1-25 ms) during the course of a syllable are
referred to as ‘gaps’ (Fig. 1).

Presentation

The presentation is identical to that used in my recent paper on Omocestus Bolivar (Ragge,
1986: 215-219). As in that study I have relied mainly on oscillograms in presenting information
on the songs and have kept the verbal descriptions as brief as possible. The references to past
accounts of the songs are not exhaustive but are intended to include all sources that the reader
might find useful to refer to for additional or confirmatory information.

The speeds of the oscillograms are (after reduction for printing) 10 mm/s, 80 mm/s, 320 mm/s
and 1280 mm/s, although the higher speeds are given only when considered to be useful. For
three species (/ineatus, festivus and stigmaticus) I have included oscillograms taken from the
calling songs of males with only one hind leg; such oscillograms, at the higher speeds, show the
individual peg-impacts with particular clarity (see Figs 30, 61, 93).

The song as a taxonomic character

In my paper on Omocestus I outlined the history of the use of the songs of Orthoptera in
taxonomy (Ragge, 1986: 219-220) and more recently (Ragge, in press) I have emphasized the
particular importance of the songs in the taxonomy and biogeography of several groups of
western European Orthoptera. The taxonomic value of the male calling songs of the western
European species of Stenobothrus can be seen at a glance from Figs 9-17. The song-patterns
show a wide variety from the long single echeme of lineatus, lasting more than 10s and with a
syllable repetition rate of less than 2/s, to the rapid series of short echemes of grammicus and
bolivarii, in which each echeme lasts less than 0-2 s and the syllable repetition rate is more than
50/s.

S. rubicundulus stands apart from the other species included in the present study in producing
a loud wing-buzz, often during hovering flight, in addition to the more usual femoro-alary
stridulation. The wing-buzz clearly forms part of this species’ communicative behaviour and
often follows immediately after a femoro-alary echeme (Fig. 17). According to Dr C. Walter,
whose observations are quoted by Harz (1975: 777), S. cotticus also produces a wing-buzz; I have
not yet been able to study the acoustic behaviour of this species, which is known only from a
small area of the French Alps.

The calling song of /ineatus is unique among all known Gomphocerine songs in having such
long syllables, each usually lasting 0-6—0-8 s. This species also has a uniquely dense stridulatory
file, so that, in spite of the slow leg movements, the peg-impact rate (for one hind leg) is still of
the order of 500/s in the middle of the downstroke (see Fig. 30). S. /ineatus is also unique among
the western European species of Stenobothrus in producing an echeme lasting more than 10s.

Among the other species in which the calling song normally consists of a single echeme
(fischeri, festivus, stigmaticus and apenninus), the song conforms to the same basic pattern of a
simple series of syllables, beginning quietly but soon reaching maximum intensity (see Figs 11,
12, 15, 16). The number and repetition rate of the syllables enable fischeri and festivus to be
distinguished from the other two species and also from each other. However, stigmaticus and
apenninus are extremely similar to each other in both calling and courtship songs; although the
echemes of stigmaticus are on average longer than those of apenninus and are composed of more

398 D. R. RAGGE

ECHEME

ae a

Lier seal re

SYLLABLE GAPS 0-05 s J

Stenobothrus stigmaticus

ECHEME-SEQUENCE

.

eS)
LOls 4

SYLLABLES
ECHEME Loos sea;

Stenobothrus grammicus

Fig1 Oscillograms of the male calling songs of two species of Stenobothrus, showing the terminology used
in this paper.

SONGS OF THE WESTERN EUROPEAN Stenobothrus 399

syllables, there is an overlap in both these song characters and I have not been able to find a
constant difference between the two songs. As these two species are completely allopatric I
think it likely that apenninus is a recent derivative from stigmaticus that has diverged from it
morphologically as a result of isolation in the Apennine mountain range. It would be most
interesting to establish experimentally whether there is any ethological barrier to intercopula-
tion between these two species.

The remaining three species, nigromaculatus, grammicus and bolivarii, have calling songs
consisting of a sequence of echemes; they also differ from the other six species in having syllable
repetition rates of more than 50/s. As can be seen from Figs 10, 13 and 14, the calling song of
nigromaculatus differs markedly from those of grammicus and bolivarii in that the constituent
echemes are far fewer in number (usually 2—6) and much longer (usually lasting 1-2 s). The very
short, rapidly repeated echemes of grammicus and bolivarii show a marked contrast with the
other seven species, but sound similar to each other to the unaided ear, differing mainly in the
syllable structure within the echemes. These two species are also similar morphologically and
may well have evolved from common stock quite recently.

As I have recently emphasized (Ragge, 1986: 220), the value of the songs as indicators of
phylogeny is very limited. Their taxonomic importance, as would be expected of a mate
recognition system, is almost entirely at the specific and subspecific levels. This can be
exemplified by some fairly close intergeneric resemblances in song between certain species of
Stenobothrus and Omocestus. The male calling song of S. apenninus, for example, is quite
similar to that of O. raymondi (Yersin) (Ragge, 1986: figs 64-72), and there is also quite a close
resemblance between the male calling songs of S. fischeri and O. broelemanni (Azam) (Ragge,
1986: figs 90-98). The members of each of these pairs are allopatric and so they have no need for
differing recognition signals (although in each case there is a marked difference in courtship
song). The calling song of all four of these species is a simple echeme and has presumably not
been subject to the selective pressures needed for further differentiation. Echemes of this basic
type also occur in several species of Chorthippus Fieber.

New lectotype designations

Gomphocerus (Stenobothrus) Bolivarii Brunner, in Bolivar, 1876: 327.
Dr A. Kaltenbach of the NM, Vienna has kindly sent me a number of syntypes of this species, and I have
selected and labelled a male as lectotype. I have labelled two other males and six females as paralectotypes.

Stenobothrus apenninus Ebner, 1915: 551.

Dr Kaltenbach has also kindly lent me six syntypes of this species, and I have selected and labelled an adult
male as lectotype; this specimen was already labelled ‘typus’ by Ebner. I have labelled the remaining five
specimens (two adult males, one adult female and two female nymphs, all labelled ‘paratypus’) as
paralectotypes.

STENOBOTHRUS Fischer

Stenobothrus Fischer, 1853: 313. Type-species: Gryllus lineatus Panzer, by subsequent designation (Kirby,
1910: 160).

Stenobothrodes Tarbinskii, 1948: 116 [as subgenus of Stenobothrus Fischer]. Type-species: Stenobothrus
eurasius Zubovskii, by monotypy. Syn. n.

Crotalacris Chopard, 1952: 280 [as subgenus of Stenobothrus Fisher]. Type-species: ‘Stenobothrus rubicun-
dus (Germar)’ [= Stenobothrus rubicundulus Kruseman & Jeekel (see p. 422)|, by monotypy. [Synony-
mized with Stenobothrodes Tarbinskii by La Greca, 1965: 70.]

Diacnosis. O'. Head of typical Gomphocerine shape, foveolae well developed. Antennae usually
filiform or slightly thickened towards tip, exceptionally clubbed (males of S. clavatus Willemse). Pronotal
lateral carinae varying from sharply incurved to almost straight. Brachypterous to macropterous. Fore
wings without bulge on anterior margin of precostal area; medial area usually widened, often conspicuous-
ly so, with Cu, and Cuz sometimes partly or even completely fused. Hind wings varying from transparent to
strongly smoky, sometimes with enlarged medial area. Ovipositor with lateral tooth on each valve.
Tympanal aperture slit-like.

400 D. R. RAGGE

Discussion. Stenobothrus can be recognized reliably only by the ovipositor, which has a lateral tooth on all
four valves. Males can sometimes be confused with Omocestus, but in most cases can be recognized by the
widened medial area of the fore wings. Both sexes of most species have somewhat broader pale bands along
the pronotal lateral carinae than are shown by Omocestus and other related genera.

The subgenera Stenobothrodes and Crotalacris are certainly not distinct from each other at the
subgeneric level, and even considered together they do not in my opinion justify recognition as a distinct
subgenus. On the other hand, I have followed Harz (1975) and other recent workers on the European
fauna in regarding Omocestus and Myrmeleotettix Bolivar as distinct genera rather than subgenera of
Stenobothrus; Stauroderus Bolivar and the Nearctic genus Bruneria McNeill, neither of which has a
toothed ovipositor, are similarly excluded.

As in the other large Gomphocerine genera Omocestus and Chorthippus, the male calling songs show a
wide range of patterns in Stenobothrus, varying from the long single echeme of lineatus to the rapid
sequence of short echemes of grammicus and bolivarii, and the remarkable wing-buzz of rubicundulus.

DistriBuTION. All Europe, the larger Mediterranean islands, North Africa and temperate Asia as far as
Mongolia.

INCLUDED SPECIES. S. amoenus (Brisout), S. apenninus Ebner, S. berberus Uvarov, S. bolivarii (Brunner),
S. bulgaricus Ramme, S. burri Karabag, S. carbonarius (Eversmann), S. caucasicus Dovnar-Zapolskii, S.
clavatus Willemse, S. cobresianus Bei-Bienko, S. cotticus Kruseman & Jeekel, S. crassipes (Charpentier),
S. croaticus Ramme, S. eurasius Zubovskii, S. festivus Bolivar, S. fischeri (Eversmann), S. graecus
Ramme, S. grammicus Cazurro, S. kirgizorum Ikonnikoy, S. lineatus (Panzer), S. maroccanus Uvarov, S.
miramae Dirsh, S. newskii Zubovskii, S. nigromaculatus (Herrich-Schaffer), S. palpalis Uvarov, S.
posthumus Ramme, S. rubicundulus Kruseman & Jeekel, S. stigmaticus (Rambur), S. sviridenkoi Ramme,
S. tadzhicus Mishchenko, S. ursulae Nadig, S. werneri Adelung, S. zubowskyi Bolivar.

Keys to the principal western European species of Stenobothrus

The only western European species not included in these keys are very local forms, sometimes of
doubtful status, in which the song is unknown to me.

Two keys are provided here, one based on morphological characters but with notes on song
differences added where appropriate, and one based entirely on the male calling songs. Some of
the species are difficult to separate morphologically and, in the absence of song information,
some specimens may not be correctly identified using the morphological key. Reference should
in any case always be made to the notes given on recognition in the accounts of each species.

Key based primarily on morphological characters

This key is for sexually mature adults, either freshly collected or showing good colour preservation. The
terminology used for wing characters is illustrated in Fig. 3.

1 Hind wings vestigial in both sexes; fore wings reduced to about the same length as the

pronotunim 9 (Apennime mountains)! .2..5. 26: sons-ce-ceew cetwecsecsutenacehetecbenestk apenninus(p. 418)
= Both pairsof wings welldeveloped : :22. 25.521. csecBegossanthateerert cs oases clan co ceaceseeenoeee eee 2
2 Palpsibiackiondankebrownat theytipieessssisecseer acer a eee eseeeee ete tee tee ee eee grammicus(p. 410)
—) ‘Palpsnotblackordark browmnlatithe tip e-n-ccasesce-co-es eee ereee eee e een eer eee cent eee eee ee eeeee 3
3 Mad ei ic2 cog atau dee ostninaateon wenaareetionictic shisha cisioeces Goa iove ee cet ae ete i ale ee eee eee Oe ee 4
= (BOM ale: oo acces ste ne ceiete ene oe dercactee ao emcl eee ete ae eee ee ont ee iceland, ae 10
4 Last segment of the maxillary palps bulbous and usually coloured orange-pink, red-brown or
Vellow DIOWily 3.....505 caakacncuiscee: Serueaas sets RES ee ae are eee erent ne pene eae bolivarii(p. 413)
— Last segment of the maxillary palps not bulbous and not coloured pink or brown ................... 5
5 Pronotal lateral carinae clearly angled in the prozona and interrupted at the angle by an
additional'transverse sulcuS(Pig. 2) sc c.asateye see eet eeteat area eer eee ee festivus(p. 410)
— Pronotal lateral carinae not or hardly angled in the prozona and not interrupted by an
additional'sulcus (mot as tm Fig.2)). che atts woe aeak she cae cee ae ase ence eee eee ee nee ee 6
6 Stigmawithin3 mnofthe fore wingtip’ ios eaccsoes. fos os seeee dees eae (eae oe es oe ee a
=» Stigma further than 3 mmifrom the fore: wing=tipyec..1.ce-se- oo, see ee ee eee oa eee ee ee 8

7 Medial area of the fore wing enlarged, as in Fig. 5, Cu, becoming fused with Cuz within the
proximal half of the fore wing. Total length more than 14 mm. (Song a sequence of two or
moreechemes, as im Pigs 50-34) en -aae- eenece eee ee eee hae ee eae nigromaculatus(p. 405)

SONGS OF THE WESTERN EUROPEAN Stenobothrus 401

medial area stigma

medial area M
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Figs 2-8 Pronotum and wings of species of Stenobothrus. 2. Dorsal view of the pronotum of S. festivus.
3. The right male fore and hind wings of S. rubicundulus, showing the principal veins and the wing
terminology used in this paper. 4—7. The right male fore wing of (4) S. lineatus, (5) S. nigromaculatus, (6)
S. fischeri, (7) S. stigmaticus. 8. The right male fore and hind wings of S. rubicundulus.

402

18

D. R. RAGGE

Medial area of the fore wing not enlarged, as in Fig. 7, Cu, remaining separate from Cu until
the distal half of the fore wing. Total length less than 14 mm. (Song almost always a single

echeme, asin Hie; 15 ex 25. ta asie to haba eeee cae eater a aa ee el ae stigmaticus(p. 413)
Cu, fused with Cuz at or near the base of the fore wing, as in Fig. 4. Cerci conical. (Song with
syllable repetition cate less than 2)S).-33i5.o-.-02es.e2te- ahaa besser meee eee ee lineatus(p. 404)

Cu, remaining distinct from Cuz in much of the proximal half of the fore wing (although the two
veins often touch in places), not as in Fig. 4. Cerci laterally compressed towards the tip.

(Song with syllable. repetition rate more that 4/s)\\ 22.2 - <2. 2d-cne)-2222eaeee Saeco sor ei'> sees 9
Fore wings very broad, as in Fig. 8. Hind wings as in Fig. 8, with the more distal part of the

anterior margin strongly sclerotized and a much enlarged medial area........ rubicundulus(p. 422)
Fore wings much narrower, asin Fig. 6. Hind wings not asin Fig. 8 .................06.06 fischeri(p. 405)
Cu, fused or continguous with Cuz in at least part of the proximal half of the fore wing............. 11
Cu, remaining separate from Cu, until the distal half of the fore wing .........................00e0eeeee 14
Stigma within 4 mnrof the fore WiING-lIp ... 6 2<1b.90n0<n neuen eine rmmalonnvacionne nigromaculatus(p. 405)
Stigma furthenthan 4 mim fromthe fore wWithp-tip 5... 2<.0c.is»caasaceseaenvoareqe-sateveesneenaes teen 12
Cu, fused with Cuz at or near the base of the fore wing ..................cseeeeeeeeeeeeeenees lineatus(p. 404)
Cu, remaining distinct from Cu in much of the proximal half of the fore wing (although the two

veins sometumes touch or fuse in places). 6 x..sacddacece sent ccete sone os ere cee acne eae ee 13
Hind wings as in Fig. 8, with the more distal part of the anterior margin strongly sclerotized and

amuch enlarged medial area card ies.... cece devoceee ve eeeseeeenc sence eee eee rubicundulus(p. 422)
Hind wingsnotas in Fig® 2 shee at hoc at aes ait bcm at eee eee fischeri(p. 405)
enethofthetorewines lessithant lima e.-eeeeecs-eeceee eee tee ee eee eee eeeee eee ree stigmaticus(p. 413)
Length of the fore wings more tham 11) mmo... 0.5 vee cans ceive saewnou the Ceaerenaceeseh eae eee 15
Hind wings as in Fig. 8, with the more distal part of the anterior margin strongly sclerotized and

amuch enlarged mediallarea. [22.5 .cis0.<2 cateoeceesns andes .eee eee eee eeee ene rubicundulus(p. 422)
Hind wings not. as i Pig. 8 «0. esas «desictgs adap ties as onin os tile sents oltre prec ae Soe eee ee 16
Eengthofithe tore wines more thane Spano ieeeen a eeeceeeteeeaeeeee ener eee eee eee ee bolivarii(p. 413)
Ienpth ofthe fore: wingsilessthamilspmunis sasnesee o-- cr cee aes eeneee nee eeetee ae eee ee 17

Pronotal lateral carinae clearly angled in the prozona and interrupted at the angle by an
additional transverse sulcus (Fig. 2). Cu; and Cuz diverging in the proximal half of the fore
WHS. aaah wines taomew saints Dales ebro cae Seats es Pea ee SD festivus(p. 410)
Pronotal lateral carinae not or hardly angled in the prozona and not normally interrupted by an
additional sulcus (not as in Fig. 2). Cu, and Cu converging or running parallel in the

proximal half of the fone: Wing 0x sos chess oS 0 «cine wince so doneslo cob one one REN see eee 18
Stigma within 4mm of the fore WiRB IP o.5..S.. 0.5- donmanaas ann ocwanniensivdninna cannes nigromaculatus(p. 405)
Stigma further than 4mm from the fore WINg-tip .....6522.-..cs0scesercosermccecersuncdeastes fischeri(p. 405)

Key based on song characters (see especially Figs 9-17)

This key is based on the calling songs produced by isolated males in warm, sunny conditions. Many of the
song differences can be easily detected by the human ear assisted by the second hand (or digital count) of a
watch, but a few of the couplets require oscillographic analysis, or at least the ability to play back the
recorded song at a slower tape speed. I have been unable to find a consistent difference between the calling
songs of stigmaticus and apenninus (couplet 9), but these species are completely allopatric and easily
separated morphologically.

See p. 397 for definitions of the terms ‘echeme’, ‘syllable’ and ‘gap’.

1

Song consisting, at least partly, of aloud buzz produced by wing-vibration, either on the ground

orin flight (Figs 109046) cosas as2.cwecSante teach cee eee eee eee teen eee ee rubicundulus(p. 422)
Song produced entirely by femoro-alary StrithulahOm 222 ooo coc cwnaem ass susioms Jovenen ao sepaaeanee 2
Song asequence of echemes in rapid succession (at least U/S) <<<. cceces eer oelsveee scent sacnatepen se Qen 3
Song a single echeme or a sequence of echemes repeated at the rate of one every 2 s or more

SLOWLY, cdosesave nsec dauctnge suena comes de tenatlplent tae neeenistaee tease me geinns tan eines oskioeing sceioelsa sets eee eae eee “4
Each echeme lasting less than 80 ms, loudest towards the end (Figs 62-72) ....... grammicus(p. 410)
Each echeme lasting more than 80 ms, loudest at the beginning (Figs 73—76) ........ bolivarii(p. 413)
Song consisting of a single echeme lasting more than 10 s and produced by very slow leg

movements (less than 2/s); syllables lasting 0-5—1-0s (Figs 18-30) .................2.. lineatus(p. 404)
Song consisting of one or more echemes, each lasting less than 10s and produced by more rapid

leg movements (at least 3/s); syllables lasting less than 0-2§ ................ceeeceeeeeeeeeeeeeeeneenes 3

Echeme usually lasting more than 5 s, with the syllables, at least in the early part of the echeme,

403

SONGS OF THE WESTERN EUROPEAN Stenobothrus

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composed of clearly separated upstroke and downstroke sounds of equal amplitude, as in

Figs £03 N06. 0220 fae nsve wacdegeunsiaes oo teak tate oe ee ee Sg eee IR a eae rubicundulus(p. 422)
— Echeme lasting less than 5 s, with the syllables composed mainly of downstroke sounds (not as
IM FIgs VOS—NOB). ek sad sees nee seca d awan eee eee eeeolabe acerca meek Re 6

6 Song consisting of at least 2 echemes; syllable repetition rate more than 50/s (Figs 31-44)
nigromaculatus(p. 405)
— Song almost always consisting of a single echeme; syllable repetition rate less than 20/s ........... fj
7 Echeme composed of more than 40 syllables and usually lasting more than 3 s (Figs 45-53)
fischeri(p. 405)

— Echeme composed of fewer than 40 syllables and usually lasting less than3s ..................-...05. 8
8 Syllable repetition rate less than 12/s; early part of echeme with about 7-8 gaps in each syllable
(Bigs SA Gil aoe: iy to ah oa oa gamete wale ees coh pe eee Soeree oene cae Mette eee festivus(p. 410)

— Syllable repetition rate usually more than 12/s; early part of echeme with fewer than 5 gaps in

each syllable (E108) 7 F102) oo ssit dost cases Suece sedel nok «ebaiiesetuncaciebamsesnaseeoeesaes Ree eae aman
9 ‘Occurring oniyin.the Apennine mountains <1... 5. ois 32a ner~ oo eee erercomeea nde eeone apenninus(p. 418)
— Notoccurringinthe Apennine mountams ... 05-2. .0.. c-ctosceeencene a sense scone seater stigmaticus(p. 413)

The songs of the western European Stenobothrus

Stenobothrus lineatus (Panzer)
(Figs 4, 18-30)

Gryllus lineatus Panzer, 1796: 9. Type(s): GERMANY (lost); the two ‘neotypes’ designated by Harz (1975:
759) are clearly invalid.

REFERENCES TO SONG. Oscillogram: Dumortier, 1963; Elsner, 1974a, 1975; Elsner & Popov, 1978; Grein,
1984; Haskell, 1957, 1958. Diagram: Bellmann, 1985a; Duijm & Kruseman, 1983; Haskell, 1957; Jacobs,
1950, 1953; Luquet, 1978; Ragge, 1965. Frequency information: Dumortier, 1963; Haskell, 1957, 1958.
Musical notation: Yersin, 1854. Verbal description only: Baier, 1930; Broughton, 1972; Chopard, 1922,
1952; Faber, 1928, 1953; Harz, 1957; Haskell, 1955. Disc recording: Andrieu & Dumortier, 1963; Grein,
1984; Ragge, Burton & Wade, 1965. Cassette recording: Bellmann, 1985b.

RECOGNITION. Males of this species may be distinguished from the rather similar species nigromaculatus
and fischeri by the cerci, which are simply conical (laterally compressed towards the tip in the other two
species). In both sexes the veins Cu; and Cuz are more completely fused together in the fore wings than in
nigromaculatus and fischeri, and the hind wings are more strongly smoky.

In the field males may be recognized at once by their highly characteristic calling song, in which the
syllable repetition rate (under 2/s) is lower than in any other European grasshopper.

SonG (Figs 18-30). The calling song of this species is unique among the western European
Gomphocerinae in both the slowness of the leg movements and the quality of the sound
produced. It consists of an echeme lasting about 10—25 s and composed of about 15-35 syllables
repeated at the rate of about 1-2—1-5/s. The echeme usually begins quietly but reaches maximum
intensity within a few syllables. The sound is usually continuous, with a highly characteristic,
rather wheezy quality, and showing slight fluctuations in intensity corresponding with the leg
movements — though partly masked by the fact that one hind leg always moves slightly in
advance of the other.

In the presence of a female the male usually produces longer echemes (with a more gradual
crescendo at the beginning), sometimes lasting a minute or more but otherwise similar to the
calling song. Between these echemes the male produces a quite different ‘ticking’ sound (Figs
21, 22, 26, 27) by moving the hind legs through a much smaller angle than in the calling song and
at a much higher rate (usually S—7 syllables/s). This ticking sound lasts typically for about half a
minute before another echeme of the calling song type is produced, but it may be greatly
prolonged, sometimes lasting for even longer than an hour. The ticking sound and the ‘calling
song’ echemes usually alternate several times before the male attempts to copulate with the
female. The courtship song of this species and the leg movements that produce it have been
described in great detail by Elsner (1974a, 1975).

DISTRIBUTION. Widespread in Europe, but not reaching further north than southern Sweden and limited to
mountains in the southern peninsulas. The range extends eastwards across central Asia to Mongolia.

SONGS OF THE WESTERN EUROPEAN Stenobothrus 405
Stenobothrus nigromaculatus (Herrich-Schaffer)
(Figs 5, 31-44)

Acridium nigromaculatum Herrich-Schaffer, 1840: 10. Type(s) unknown; the two ‘neotypes’ designated by
Harz (1975: 761) are clearly invalid.

REFERENCES TO SONG. Oscillogram: Grein, 1984; Schmidt & Schach, 1978. Diagram: Bellmann, 1985a;
Duijm & Kruseman, 1983; Jacobs, 1950, 1953. Verbal description only: Faber, 1928, 1953; Harz, 1957. Disc
recording: Grein, 1984. Cassette recording: Bellmann, 1985b.

RECOGNITION. Males of this species may be distinguished from lineatus by the cerci, which are laterally
compressed towards the tip (simply conical in Jineatus). In the fore wings of both sexes the veins Cu, and
Cuz are less completely fused together than in /ineatus, and the stigma is nearer the wing-tip than in fischeri
(cf. Figs 5, 6).

In the field males may be recognized quite easily by their calling song, in which the syllable repetition rate
(about 70—110/s) is much higher than in /ineatus (about 1-2—1-5/s) and fischeri (about 10—15/s).

Sone (Figs 31-44). The calling song consists of a series of 2-6 echemes (each beginning quietly
but showing a rapid crescendo), separated by intervals of 1-3 s. Each echeme lasts about 0-8—2-0
s (rarely up to 2-5 s) and consists of about 90—200 syllables (rarely up to 250) repeated at the rate
of about 70—110/s.

In the presence of a female the male produces a quite complicated courtship song, consisting
of two alternating phases (Figs 35, 36, 41-44). The first phase consists of a series of fairly quiet
echemes separated by intervals of about 0-6—0-9 s; during these intervals a few even quieter
sounds are often produced. Each echeme lasts for about 0-3—0-9 s and is composed of about
40-80 syllables repeated at the rate of about 80—100/s. This phase lasts for an indefinite period,
usually more than 30 s and often several minutes. The second phase, which follows abruptly,
consists of a series of about 3-7 louder and longer echemes, each of which is preceded by a much
shorter echeme. The shorter echemes last for less than half a second, and in fact after the first
longer echeme they are often reduced to little more than a single syllable. The longer echemes
usually last for about 0-8—2-2 s and are subdivided into two parts, one with slower syllables
(about 40-—60/s) and one with faster syllables (about 70—120/s). At the end of this second phase
the male either attempts to copulate with the female or, after a short pause, begins the first phase
again.

Note that the difference in temporal pattern between the oscillograms taken from recording
482/3 (Figs 35, 41, 42) and those taken from recording 184/1 (Figs 36, 43, 44) may have been
caused, at least partly, by the fact that recording 184/1, unlike any other field recording used in
this study, was made in conditions of hazy sunshine, so that the subject was receiving less radiant
heat than it would have in full sunshine.

DIsTRIBUTION. Southern Europe and central Asia; absent from the Iberian Peninsula except in the vicinity
of the Pyrenees, and confined to uplands in the Italian and Balkan Peninsulas.

Stenobothrus fischeri (Eversmann)
(Figs 6, 45-53)

Oedipoda fischeri Eversmann, 1848: 11. Syntypes of both sexes, U.S.S.R.: Ural Mountains (ZI, Lenin-
grad) [1 o’ , 1 2 syntypes examined].

REFERENCES TO SONG. Diagram: Luquet, 1978. Verbal description only: Faber, 1953; Harz, 1957.

RECOGNITION. Males of this species may be distinguished from /ineatus by the cerci, which are laterally
compressed towards the tip (simply conical in /ineatus). In the fore wings of both sexes the veins Cu, and
Cuz are less completely fused together than in Jineatus and the stigma is further from the wing-tip than in
nigromaculatus (cf. Figs 5, 6).

In the field the single echeme of the male calling song, which lasts for 3—5 s and is composed of more than
40 syllables, enables this species to be distinguished from Jineatus, nigromaculatus, grammicus and
bolivarii. The calling songs of festivus and stigmaticus are more similar, but in both these species the
echeme is shorter, with fewer than 40 syllables.

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SonG (Figs 45-53). The calling song is a single echeme lasting about 3-5 s and consisting of about
40-55 syllables repeated at the rate of about 12—13/s. The echeme begins quietly, usually
reaching maximum intensity about halfway through its duration. Oscillographic analysis shows
that there are gaps in each syllable. Previously published accounts of the calling song of this
species have been based almost entirely on Faber (1953), who gave lower figures for both the
duration of the echeme (‘2 oder etwas mehr Sekunden’) and the syllable repetition rate
(8-914/s), although suggesting that these figures applied to males singing in full sunshine.
Faber’s observations were made in the laboratory on males collected for him from Oberweiden,
east of Vienna in Lower Austria, whereas mine were made in the field in southern France; in
France I would expect such a low syllable repetition rate only in dull weather or at unusually low
air temperatures.

In the presence of a female the male produces a long and complicated courtship song (Figs 47,
50, 53), which has been fully described by Faber (1953: Beilage 4).

DISTRIBUTION. In western Europe this species seems to be confined to two isolated areas: southern France
and central Spain. Further east it occurs from Austria eastwards across eastern Europe (including the
Balkan Peninsula) and central Asia to Mongolia.

Stenobothrus festivus Bolivar
(Figs 2, 54-61)

Stenobothrus festivus Bolivar, 1887: 94. Syntypes of both sexes, Spain: Uclés, Aranjuez, San Ildefonso,
Escorial, Gredos, Navacerrada (7MNCN, Madrid).

REFERENCES TO SONG. No published work known to me.

RECOGNITION. This species is perhaps most likely to be confused with grammicus or bolivarii, which share
with festivus the strongly inflexed pronotal lateral carinae. However, males of festivus lack the modified or
characteristically coloured last palpal segments shown by the other two species, and in both sexes the
pronotal lateral carinae are more sharply angled in the prozona, where they are clearly interrupted by an
additional transverse sulcus on each side of the pronotum (Fig. 2).

In the field the single echeme of the male calling song enables this species to be distinguished easily from
grammicus and bolivarii, which both produce a rapid sequence of echemes. The male calling songs of
fischeri and stigmaticus are more similar, but fischeri has a longer echeme, with more than 40 syllables
(usually fewer than 30 in festivus) and in stigmaticus the syllable repetition rate is usually more than 12/s
(less than 12/s in festivus).

Sonc (Figs 54—61). The calling song is a single echeme lasting about 2—3 s and consisting of about
20-30 syllables repeated at the rate of about 7—11/s. The echeme usually begins quietly, reaching
maximum intensity about halfway through its duration or a little earlier, but sometimes it begins
more abruptly and sometimes there are one or more louder syllables before the quiet beginning.
Oscillographic analysis shows that there is a characteristic pattern of gaps in the syllables: at the
beginning of the echeme the syllables are completely broken up by gaps, but as the echeme
progresses the gaps gradually disappear so that there are usually no gaps in the last few syllables.
In a song produced by only one hind leg the gaps often persist until rather later in the echeme
(Fig. 57), since there is no tendency for the gaps in the syllables produced by one hind leg to be
obscured by sounds produced by the other.

DISTRIBUTION. Widespread in the Iberian Peninsula and occurring locally in southern France.

Stenobothrus grammicus Cazurro
(Figs 62-72)

Stenobothrus grammicus Cazurro, 1888: 457. Syntypes of both sexes, SPAIN: Navacerrada, Penalara,
Escorial, San Ildefonso, Ofia (7MNCN, Madrid).

REFERENCES TO SONG. Diagram: Luquet, 1978.

RECOGNITION. Both sexes of this species may be recognized easily by the palps, which are black or dark

411

SONGS OF THE WESTERN EUROPEAN Stenobothrus

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SONGS OF THE WESTERN EUROPEAN Stenobothrus 413

brown at the tip. In addition the male antennae are curved outwards and noticeably thickened and
darkened towards the tip.

In the field the distinctive calling song, consisting of about 10-30 very brief and rapidly repeated
echemes, enables males to be easily distinguished from most other species of Stenobothrus; however,
oscillographic analysis is necessary to show clearly the difference from the calling song of bolivarii, which
has longer echemes with a rather different syllable structure (see below).

Sonc (Figs 62—72). The calling song consists of a sequence of about 10-30 echemes lasting about
5-15 s; the echemes are thus repeated at the rate of about 2/s. Each echeme lasts about 40-80 ms
and is composed of a series of 4—8 syllables. The only other species of Stenobothrus known at
present to have a rather similar calling song is bolivarii, but oscillographic analysis shows that
species to have longer echemes, lasting about 80-110 ms, with a rather different syllable
structure: in grammicus the loudest syllables are towards the end of each echeme, whereas in
bolivarii the loudest syllable is at the beginning (cf. Figs 71, 76).

Luquet (1978: 427) gives the rather lower number of 6—8 echemes in each sequence of the
calling song, based on observations in the Mont Ventoux area of Vaucluse in SE. France. My
own studies of four males at two localities, also in the region of Mont Ventoux, suggest that this
number is unusually low.

In the presence of a female the male produces a series of quieter and much longer echemes
(Figs 64, 67, 70). Each of these echemes lasts for about 1—2 s and consists of a soft, continuous
and rather amorphous sound. Between the echemes there are intervals of varying duration
(usually between 3 and 15 s), during which quieter sounds are often produced.

DisTRIBUTION. Known only from upland areas in southern France and the Iberian Peninsula.

Stenobothrus bolivarii (Brunner)
(Figs 73-76)

Gomphocerus (Stenobothrus) bolivarii Brunner, in Bolivar, 1876: 327. LECTOTYPE: ©’, Spain: Sierra de
Guadarrama, La Granja (NM, Vienna), here designated (see p. 399) [examined].

REFERENCES TO SONG. No published work known to me.

RECOGNITION. Males of this species may be recognized by the maxillary palps, in which the last segment is
bulbous and usually coloured orange-pink or reddish brown. Females are more difficult to recognize, as the
last segment of the maxillary palps is more normal in shape and is not always distinctively coloured like that
of the male; however, in the upland areas of the Iberian Peninsula where bolivarii occurs the only other
species of Stenobothrus with which females are likely to be confused is grammicus, in which both pairs of
palps have black or dark brown tips in both sexes.

In the field the short, rapidly repeated echemes of the calling song enable males to be easily distinguished
from most other species of Stenobothrus; the calling song of grammicus is, however, rather similar and
oscillographic analysis is necessary to show clearly that bolivarii has longer echemes with a different
syllable structure.

SonG (Figs 73-76). The calling song is quite similar to that of grammicus, consisting of a
sequence of about 10—20 echemes lasting about 4—8 s; the echemes are repeated at the rate of
about 2—3/s. Each echeme lasts about 80-150 ms and is composed of a series of about 5-10
syllables; the echemes are thus rather longer than those of grammicus and show a different
syllable structure, with the loudest sound at the beginning of each echeme (cf. Figs 71, 76).

DisTRIBUTION. Known only from upland areas in the Iberian Peninsula.

Stenobothrus stigmaticus (Rambur)
(Figs 7, 77-93)

Gryllus stigmaticus Rambur, [1838]: 93. Syntypes of both sexes, Spain: Sierra Nevada (lost); the two
‘neotypes’ designated by Harz (1975: 768) are clearly invalid.

REFERENCES TO SONG. Oscillogram: Grein, 1984. Diagram: Bellmann, 1985a; Duijm & Kruseman, 1983;

D. R. RAGGE

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Jacobs, 1950, 1953; Ragge, 1965. Verbal description only: Broughton, 1972; Faber, 1928, 1953; Harz, 1957.
Disc recording: Grein, 1984; Ragge, Burton & Wade, 1965. Cassette recording: Bellmann, 1985b.

RECOGNITION. S. stigmaticus is conspicuously smaller than any other fully winged species of Stenobothrus
occurring in western Europe. Were it not for the size difference it could be confused with nigromaculatus,
but the veins Cu, and Cu, are much more widely separated in the fore wings than in that species.

In the field the male calling song, usually consisting of a single echeme lasting 1—3 s, enables this species
to be distinguished easily from all other species of Stenobothrus with overlapping distributions except
fischeri and festivus; these species produce single echemes of a similar kind to stigmaticus, but in fischeri the
echeme is longer, with more than 40 syllables (fewer than 40 in stigmaticus), and in festivus the syllable
repetition rate is less than 12/s (usually more than 12/s in stigmaticus). Both the calling and courtship songs
are extremely similar to those of apenninus, but as that species is confined to the Apennines, where
stigmaticus does not occur, confusion between the songs of these two species is unlikely. (See also the
remarks on pp. 397, 399.)

SonG (Figs 77-93). The calling song is a rather quiet echeme lasting about 1-3 s and consisting of
about 25-40 syllables repeated at the rate of about 10—20/s (usually 14-18/s). The echeme
usually begins quietly, reaching maximum intensity within about one second. Oscillographic
analysis shows that there are usually gaps in the syllables, especially near the beginning of the
echeme, but they are not always very clear. The echemes are usually produced singly and
repeated at irregular intervals, but occasionally two echemes are produced within a few seconds
of each other (Fig. 79).

In the presence of a female the male produces a series of echemes separated by intervals of
about 2-7 s (Figs 81, 86, 91). Each echeme is similar to that of the calling song, but is quieter,
longer (usually lasting 3—S s and composed of about 50-70 syllables) and with one hind leg
moving more vigorously than the other; the last few syllables are rather modified and then the
hind leg that has been moving more vigorously produces a short, loud syllable by a very quick
downstroke. The more vigorously moving hind leg alternates from one side to the other in
successive echemes. Sometimes the loud syllable at the end of each echeme is omitted.

DIsTRIBUTION. Widespread in western and central Europe, but not reaching as far north as Scandinavia. In
southern Europe it occurs in the mountainous parts of the Iberian Peninsula and the northern part of the
Balkan Peninsula but is apparently absent from the Italian Peninsula (although occurring in the Italian
Alps). It is recorded from the Rif mountains in Morocco and its range extends eastwards into European
Russia and Asia Minor.

Stenobothrus apenninus Ebner
(Figs 94-102)
Stenobothrus apenninus Ebner, 1915: 551. LECTOTYPE ©, Ivaty: Lazio, Monti Reatini,
Terminillo, 6.viii.1912 (R. Ebner) (NM, Vienna), here designated (see p. 399) [examined].
REFERENCES TO SONG. The only published reference to the song known to me is that of Harz (1975: 770), in
which he quotes Faber as having told him in 1971 that the song was indistinguishable from that of
stigmaticus.

RECOGNITION. This montane species is endemic to the Apennine range in the Italian Peninsula, where it
may be easily distinguished from other species of Stenobothrus by the vestigial hind wings of both sexes and
the much reduced fore wings of the female. In the field the calling song of the males, consisting of a single
echeme lasting 1-2 s, also enables this species to be distinguished easily from Italian species of the genus
occurring in the Apennines.

SonG (Figs 94-102). The calling song is a single echeme lasting about 1—2 s and consisting of
about 20-30 syllables repeated at the rate of about 15—20/s. The echeme usually begins quietly
but reaches maximum intensity within 0-5 s. Oscillographic analysis shows that there are clear
gaps in the syllables throughout the echeme.

In the presence of a female the male produces a quieter courtship song (Figs 96, 99, 102) very
similar to that of stigmaticus (see above) but with shorter echemes (often lasting about 2 s and
composed of about 30 syllables).

DISTRIBUTION. Known only from the Apennine range in Italy.

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Stenobothrus rubicundulus Kruseman & Jeekel
(Figs 8, 103-116)

Gryllus miniatus Charpentier, 1825: 155. Syntypes of both sexes, Austria: Alps, and GERMANY: ‘in
Saxoniae montibus’ (lost); the two ‘neotypes’ designated by Harz (1975: 775) are clearly invalid.
[Primary homonym of Gryllus miniatus Pallas, 1771: 467.]

[‘Stenobothrus rubicundus (Germar)’; Yakobson & Bianki, 1902: 223, and other authors. Misinterpreta-
tion of Germar (1817: 256) and misidentification of Gryllus (Locusta) rubicundus Goeze, 1778: 108.]
Stenobothrus rubicundulus Kruseman & Jeekel, 1967: 79. [Replacement name for Gryllus miniatus

Charpentier. |

REFERENCES TO SONG. Oscillogram: Elsner, 1974b. Verbal description only: Chopard, 1952; Faber, 1953;
Harz, 1957.

RECOGNITION. The very broad fore wings, with unbranched M, regular ladder-like cross-veins in the medial
area, and Cu, and Cu, quite clearly separated (Fig. 8), enable males of this species to be recognized quite
easily. In the female these characters of the fore wing are not so clearly developed, but the hind wings of
both sexes have an unbranched M and greatly enlarged medial area, and the more distal part of the anterior
margin is heavily sclerotized and conspicuously dark in colour (Fig. 8).

In the field males may be easily recognized by their highly characteristic calling song, which includes a
loud buzzing sound produced by wing-vibration, either on the ground or in hovering flight.

Sonc (Figs 103-116). The calling song is composed of two quite different kinds of sound: one is
produced by the usual femoro-alary stridulation and the other by wing-vibration, either on the
ground or in flight. The song sometimes consists of femoro-alary stridulation immediately
followed by wing-vibration, but each kind of sound production may occur without the other.

The femoro-alary stridulation (Figs 103-108) usually begins with a series of syllables in which
the upstroke and downstroke of the hind legs produce clearly separated, simple sounds (Figs
105, 107); the syllable repetition rate is variable, but is usually 8—13/s. After a few seconds the
pattern is often complicated by some of the downstrokes being prolonged and broken up by gaps
(Figs 106, 108). This femoro-alary echeme usually lasts for about 6-10 s, after which it may be
followed immediately by wing-vibration (Fig. 104).

The wing-vibration produces a loud buzzing sound (Figs 109-116) usually lasting about 4-10
s. According to Elsner (19745) the sound is produced by the strongly sclerotized anterior
margins of the two hind wings being beaten against each other. Oscillographic analysis shows the
buzzing to consists of a series of sharply distinct short-lived sounds repeated at the rate of about
55-85/s (Figs 111-116). The whole of the wing-buzz may be produced while the male is sitting on
the ground, but often the male takes off and continues to produce the buzzing sound in flight,
often hovering above the ground (Fig. 110, second wing-buzz).

In the presence of a female the male begins with femoro-alary syllables similar in kind to those
of the calling song, but the echeme is often greatly prolonged, sometimes lasting for more than
30 minutes. This is followed by wing-vibration and there is then often an alternation of shorter
echemes of femoro-alary stridulation and bursts of wing-vibration. A sequence of this kind may
end in the male taking off during a burst of wing-vibration and finally an attempt to copulate with
the female.

DIsTRIBUTION. This montane species is known only from the Alps, higher Apennines and mountains of the
Balkan Peninsula.

References

Andrieu, A.-J. & Dumortier, B. 1963. Chantes d’insectes. 30 cm disc, 33 rpm. Neuilly (Pacific).

Baier, L. J. 1930. Contribution to the physiology of the stridulation and hearing of insects. Zoologische
Jahrbiicher (Zoologie und Physiologie) 47: 151-248.

Bei-Bienko, G. Ya. & Mishchenko, L. L. 1951. Locusts and grasshoppers of the U.S.S.R. and adjacent
countries. Part II. Opredeliteli po Faune SSR, Izdavaemye Zoologicheskim Muzeem Akademii Nauk,
Leningrad 40: 381-667. [In Russian. English translation published by Israel Program for Scientific
Translations, Jerusalem, 1964, iv + 291 + xxi pp.]

Bellmann, H. 1985a. Heuschrecken. Beobachten, bestimmen. 210 pp. Melsungen.

SONGS OF THE WESTERN EUROPEAN Stenobothrus 423

— 1985b. Die Stimmen der Heimischen Heuschrecken. Tape cassette. Melsungen (Neumann-
Neudamm).

Bolivar, I. 1876. Sindpsis de los ort6pteros de Espana y Portugal. Segunda parte. Anales de la Sociedad
Espanola de Historia Natural 5: 259-372.

— 1887. Especies nuevas 6 criticas de ort6pteros. Anales de la Sociedad Espanola de Historia Natural 16:
90-114.

Broughton, W. B. 1972. The grasshopper and the taxonomer. III. Keys to the species. Journal of Biological
Education 6: 385-395.

Cazurro y Ruiz, M. 1888. Enumeracién de los Ort6pteros de Espana y Portugal. Anales de la Sociedad
Espanola de Historia Natural 17: 435-513.

Charpentier, T. de 1825. Horae entomologicae, adjectis tabulis novem coloratis. xvi + 262 pp. Bratislava.

Chopard, L. 1922. Orthoptéres et Dermaptéres. Faune de France 3, vi + 212 pp.

— 1952. Orthoptéroides. Faune de France 56 (1951), 359 pp.

Duijm, M. & Kruseman, G. 1983. De Krekels en Sprinkhanen in de Benelux. 186 pp. Amsterdam.

Dumortier, B. 1963. The physical characteristics of sound emissions in Arthropoda (pp. 346-373). In
Busnel, R.-G. [Ed.], Acoustic behaviour of Animals. xx + 933 pp. Amsterdam, London & New York.

Ebner, R. 1915. Zur Kenntnis der Orthopterenfauna der Abruzzen. Deutsche Entomologische Zeitschrift
1915: 545-570.

Elsner, N. 1974a. Neuroethology of sound production in gomphocerine grasshoppers (Orthoptera:
Acrididae). I. Song patterns and stridulatory movements. Journal of Comparative Physiology 88:
67-102.

— 1974b. Neural economy: bifunctional muscles and common central pattern elements in leg and wing
stridulation of the grasshopper Stenobothrus rubicundus Germ. (Orthoptera: Acrididae). Journal of
Comparative Physiology 89: 227-236.

— 1975. Neuroethology of sound production in gomphocerine grasshoppers (Orthoptera: Acrididae).
II. Neuromuscular activity underlying stridulation. Journal of Comparative Physiology 97: 291-322.
Elsner, N. & Popov, A. V. 1978. Neuroethology of acoustic communication. Advances in Insect Physiology

13: 229-355.

Eversmann, E. 1848. Additamenta quaedam levia ad Fischeri de Waldheim celeberrimi Orthoptera Rossica.
15 pp. Moscow.

Faber, A. 1928. Die Bestimmung der deutschen Geradfliiger (Orthopteren) nach ihren Lautausserungen.
Zeitschrift fiir Wissenschaftliche Insektenbiologie 23: 209-234.

— 1953. Laut- und Gebdardensprache bei Insekten. Orthoptera (Geradfliigler). Teil 1. 198 pp. Stuttgart.

Fischer, L. H. 1853. Orthoptera Europaea. xx + 454 pp. Leipzig.

Germar, E. F. 1817. Reise nach Dalmatien und in das Gebiet von Ragusa. xii + 323 pp. Leipzig &
Altenburg.

Goeze, J. A. E. 1778. Entomologische Beytrage zu des Ritter Linné zwolften Ausgabe des Natursystems 2,
Ixxii + 352 pp. Leipzig.

Grein, G. 1984. Gesdnge der heimischen Heuschrecken. Akustisch-optische Bestimmungshilfe. 30 cm disc,
33 rpm [with oscillograms]. Hannover (Niedersachsisches Landesverwaltungsamt).

Harz, K. 1957. Die Geradfliigler Mitteleuropas. xxiv + 494 pp. Jena.

—— 1975. Die Orthopteren Europas II. Series Entomologica 11, [viii +] 939 pp.

Haskell, P. T. 1955. Intensité sonore des stridulations de quelques Orthoptéres britanniques [pp.
154-167]. Jn Busnel, R.-G. [Ed.], Colloque sur l’Acoustique des Orthoptéres. Annales des Epiphyties
(fascicule spécial de 1954), 448 pp.

— 1957. Stridulation and associated behaviour in certain Orthoptera. 1. Analysis of the stridulation of,
and behaviour between, males. British Journal of Animal Behaviour 5: 139-148.

— 1958. Stridulation and associated behaviour in certain Orthoptera. 2. Stridulation of females and their
behaviour with males. Animal Behaviour 6: 27-42.

Herrich-Schaffer, [G. A. W.] 1840. Nomenclator entomologicus. Verzeichniss der europdischen Insecten
zur Erleichterung des Tauschverkehrs mit Preisen versehen 2, viii + 244 pp. Regensburg.

Jacobs, W. 1950. Vergleichende Verhaltensstudien an Feldheuschrecken. Zeitschrift fiir Tierpsychologie
7: 169-216.

—— 1953. Verhaltensbiologische Studien an Feldheuschrecken. Zeitschrift fiir Tierpsychologie, Beiheft 1,
vii + 228 pp.

Kirby, W. F. 1910. A synonymic catalogue of Orthoptera. Vol. III. Orthoptera Saltatoria. Part II.
(Locustid@ vel Acridiide). x + 674 pp. London.

Kruseman, G. & Jeekel, C. A. W. 1967. Stenobothrus rubicundus (Germar, 1817): an invalid name
(Orthoptera). Entomologische Berichten, Amsterdam 27: 78-80.

424 D. R. RAGGE

La Greca, M. 1965. Su alcuni Blattodei, Mantodei ed Ortotteri della fauna italiana, nuovo o poco noti.
Fragmenta Entomologica 4: 64-71.

Luquet, G. C. 1978. La systématique des Acridiens Gomphocerinae du Mont Ventoux (Vaucluse) abordée
par le biais du comportement acoustique [Orthoptera, Acrididae]. Annales de la Société Entomologique
de France (N.S.) 14: 415-450.

Pallas, P. S. 1771. Reise durch verschiedene Provinzen des russischen Reichs 1, [viii +] 504 pp. St.
Petersburg.

Panzer, G. W. F. 1796. Faunae insectorum Germanicae initia oder Deutschlands Insecten. Heft 33. 24 pp.
Nurnberg.

Ragge, D. R. 1965. Grasshoppers, crickets and cockroaches of the British Isles. xii + 299 pp. London & New
York.

—— 1986. The songs of the western European grasshoppers of the genus Omocestus in relation to their
taxonomy (Orthoptera: Acrididae). Bulletin of the British Museum (Natural History) (Entomology) 53:
213-249.

—— In press. Speciation and biogeography of some southern European Orthoptera, as revealed by their
songs.

Ragge, D. R., Burton, J. F. & Wade, G. F. 1965. Songs of the British grasshoppers and crickets. 17 cm disc,
33 rpm. London (Warne).

Rambur, P. 1838-1842. Faune entomologique de l’Andalousie 2, 336 pp. Paris.

Schmidt, G. H. & Schach, G. 1978. Biotopmassige Verteilung, Vergesellschaftung und Stridulation der
Saltatorien in der Umgebung des Neusiedlersees. Zoologische Beitrage (N.F.) 24: 201-308.

Tarbinskii, S. P. 1948. Order Saltatoria (Orthoptera) — jumping orthopterans [pp. 76-127]. In Tarbinskii,
S. P. & Pavilshchikov, N. N. [Eds], Keys to the insects of the European USSR. 1128 pp. Moscow &
Leningrad. [In Russian. ]

Tsyplenkov, E. P. 1970. Harmful Acridoidea of the U.S.S.R. 272 pp. Leningrad. [In Russian. English
translation published by Amerind Publishing Co., New Delhi, 1978, viii + 208 pp.]

Yakobson, G. G. & Bianki, V. L. 1902-1905. Orthoptera and Pseudoneuroptera of the Russian Empire and
neighbouring countries. x + 952 pp. Leningrad. [In Russian. ]

Yersin, [A.] 1852. Mémoire sur quelques faits relatifs a la stridulation des Orthopteéres et a leur distribution
géographique en Europe. Bulletins des Séances de la Société Vaudoise des Sciences Naturelles 4: 108-128.

Index

Invalid names are in italics; principal page references in bold.

amoenus 400
apenninus 394, 397, 399, 400,
404, 418

berberus 400

bolivarii 394, 397, 399, 400,
402, 405, 410, 413

broelemanni 399

Bruneria 400

bulgaricus 400

burri 400

carbonarius 400
caucasicus 400
Chorthippus 399, 400
clavatus 399, 400
cobresianus 400
cotticus 397, 400
crassipes 400
croaticus 400
Crotalacris 399, 400

eurasius 399, 400

festivus 394, 397, 400, 402,
404, 405, 410, 418

fischeri 393, 394, 397, 399,
400, 402, 404, 405, 410, 418

graecus 400
grammicus 394, 397, 399, 400,
402, 405, 410, 413

kirgizorum 400

lineatus 393, 397, 400, 402,
404, 405

maroccanus 400
miniatus 422
miramae 400
Myrmeleotettix 400

newskii 400
nigromaculatus 393, 399, 400,
402, 404, 405, 418

Omocestus 397, 399, 400

palpalis 400
posthumus 400

raymondi 399

rubicundulus 394, 399, 400,
402, 404, 422

rubicundus 399, 422

Stauroderus 400

Stenobothrodes 399, 400

stigmaticus 394, 397, 399, 400,
402, 404, 405, 410, 413, 418

sviridenkoi 400

tadzhicus 400

ursulae 400

werneri 400

zubowskyi 400

British Museum (Natural History)

Milkweed butterflies: their cladistics and biology
P.R. Ackery & R. I. Vane-Wright

The Danainae, a subfamily of the Nymphalidae, contains only some 150 species, yet aspects of
their biology have stimulated far more attention than can be justified by species numbers
alone. In recent years, an expansive literature has grown, considering aspects of their
courtship and pre-courtship behaviour, migration, larval hostplant associations, mimicry and
genetics. The popularity of danaines among biologists can certainly be attributed to this
combination, within one small group, of so many of the factors that make butterflies such an
interesting group to study. The obvious need to place this wealth of biological data within an
acceptable systematic framework provided theimpetus for this volume.

Started in 1976 within the conventions of evolution by natural selection and Hennig’s
phylogenetic systematics, the book is now largely about natural history (what the animals have
and do, where they live and how they develop) and natural groups — as revealed by a form of
analysis approaching that practised by the new school of ‘transformed cladistics’. The authors
have prepared a handbook that will appeal to a wide range of biologists, from museum
taxonomists to field ecologists.

425 pp, 12 pp colour, 73 b/w plates, line and graphic illustrations, maps, extensive bibliography.
ISBN 0 565 00893 5. 1984. Price £50.

Titles published in Volume 55

A reclassification of the European Tetrastichinae (Hymenoptera: Eulophidae), with a revision of
certain genera
By M. W. R. de V. Graham

The songs of the western European grasshoppers of the genus Stenobothrus in relation to their
taxonomy (Orthoptera: Acrididae)
By D. R. Ragge

Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk
Printed in Great Britain by Henry Ling Ltd, Dorchester